Anxiety and Decision-Making
2012; Elsevier BV; Volume: 72; Issue: 2 Linguagem: Inglês
10.1016/j.biopsych.2011.12.027
ISSN1873-2402
AutoresCatherine A. Hartley, Elizabeth A. Phelps,
Tópico(s)Decision-Making and Behavioral Economics
ResumoAlthough the everyday decision-making of clinically anxious individuals is clearly influenced by their excessive fear and worry, the relationship between anxiety and decision-making remains relatively unexplored in neuroeconomic studies. In this review, we attempt to explore the role of anxiety in decision-making with a neuroeconomic approach. We first review the neural systems mediating fear and anxiety, which overlap with a network of brain regions implicated in studies of economic decision-making. We then discuss the potential influence of cognitive biases associated with anxiety upon economic choice, focusing on a set of decision-making biases involving choice in the face of potential aversive outcomes. We propose that the neural circuitry supporting fear learning and regulation may mediate the influence of anxiety upon choice and suggest that techniques for altering fear and anxiety may also change decisions. Although the everyday decision-making of clinically anxious individuals is clearly influenced by their excessive fear and worry, the relationship between anxiety and decision-making remains relatively unexplored in neuroeconomic studies. In this review, we attempt to explore the role of anxiety in decision-making with a neuroeconomic approach. We first review the neural systems mediating fear and anxiety, which overlap with a network of brain regions implicated in studies of economic decision-making. We then discuss the potential influence of cognitive biases associated with anxiety upon economic choice, focusing on a set of decision-making biases involving choice in the face of potential aversive outcomes. We propose that the neural circuitry supporting fear learning and regulation may mediate the influence of anxiety upon choice and suggest that techniques for altering fear and anxiety may also change decisions. The tendency to experience anxiety is a relatively consistent individual trait (1Spielberger C.D. Anxiety: Current Trends in Theory and Research: I. Academic Press, Oxford, England1972Google Scholar), suggesting that it has stable underlying neural substrates and may be an important factor driving behavioral variation in a variety of domains, including decision-making. In patients suffering from anxiety disorders, heightened anxiety interferes with the ability to adaptively function in everyday tasks, such as employment or social relations. Although it is clear for these patients that their pathological anxiety influences their daily decisions, a more nuanced understanding of the relationship between anxiety and decision-making is needed. Although anxiety has long been known to involve behavioral aberrations in the face of potential negative outcomes, the burgeoning field of neuroeconomics provides a structured approach to studying the computational and neurobiological mechanisms underlying this dysfunction. Neuroeconomic studies typically define mathematically the optimal or normative behavior in a decision-making task, allowing precise quantification of individual deviations from these norms. These parameters can then be used to probe the neural correlates of decision biases. Characterizing the specific decision biases occurring with anxiety may enhance our understanding of the consequences of individual variability in nonclinical trait anxiety as well as the nature of the dysfunction underlying anxiety disorders. In this review, we build on behavioral and neuroeconomic principles of decision-making to explore the impact of anxiety on specific decision variables. We first review the neural systems implicated in fear and anxiety, which overlap with those highlighted in neuroeconomic studies of decision-making. We then discuss how anxiety-induced alterations in the brain circuitry of fear result in predictable cognitive biases that may influence later choices. We review the relatively few studies using behavioral economic paradigms to characterize how anxiety influences decision variables, speculating as to how the effects of anxiety on choice may arise from cognitive biases associated with anxiety and the neural circuitry implicated in fear. Finally, we discuss how techniques for altering fear and anxiety may also change decisions. Fear and anxiety share many common cognitive and physiological properties; however, they can also be distinguished (2Sylvers P. Laprarie J. Lilienfeld S. Differences between trait fear and trait anxiety: Implications for psychopathology.Clin Psychol Rev. 2011; 31: 122-137Crossref PubMed Scopus (242) Google Scholar). Fear responses are elicited by specific stimuli and tend to be short-lived, decreasing once a threat has dissipated. Anxiety may be experienced in the absence of a direct physical threat and typically persists over a longer period of time. However, anxiety is commonly conceptualized as a state of sustained fear (3Davis M. Walker D.L. Miles L. Grillon C. Phasic vs sustained fear in rats and humans: role of the extended amygdala in fear vs anxiety.Neuropsychopharmacology. 2010; 35: 105-135Crossref PubMed Scopus (1069) Google Scholar). Studies of fear neurocircuitry highlight a network of brain regions enabling the adaptive expression of fear to potential threats and its inhibition and control with safety. Details of this circuitry have been investigated across species using classical fear conditioning as a model paradigm. During fear conditioning, a previously neutral stimulus, such as a tone, is paired with an intrinsically aversive stimulus, such as an electric shock, eliciting a range of automatic, unconditioned fear responses (4Davis M. The role of the amygdala in conditioned and unconditioned fear and anxiety.in: Aggleton J.P. The Amygdala. Oxford University Press, Oxford, UK2000: 213-288Google Scholar, 5LeDoux J.E. The amygdala.Curr Biol. 2007; 17: 868-874Abstract Full Text Full Text PDF PubMed Scopus (1031) Google Scholar). After one or more tone-shock pairings, presentation of the tone alone is sufficient to elicit a fear response, the conditioned response, providing evidence of a learned association between the two stimuli. Once acquired, conditioned fear can be diminished via a number of techniques (6Hartley C.A. Phelps E.A. Changing fear: The neurocircuitry of emotion regulation.Neuropsychopharmacology. 2010; 35: 136-146Crossref PubMed Scopus (369) Google Scholar). In extinction, the tone is presented repeatedly without the shock, resulting in a gradual decrease in conditioned fear expression. Evidence that fear can return after successful extinction training after the passage of time (spontaneous recovery), changes in context (renewal), or stress (reinstatement) suggests the original fear memory is not erased with extinction but is rather inhibited during the expression of extinction learning (7Bouton M.E. Context and behavioral processes in extinction.Learn Mem. 2004; 11: 485-494Crossref PubMed Scopus (1394) Google Scholar). In humans, intentional cognitive strategies including emotional suppression, redeployment of attention, or cognitive reinterpretation or reappraisal of the significance of a stimulus may also be used to diminish fear (6Hartley C.A. Phelps E.A. Changing fear: The neurocircuitry of emotion regulation.Neuropsychopharmacology. 2010; 35: 136-146Crossref PubMed Scopus (369) Google Scholar). The neurocircuitry supporting fear conditioning has been extensively investigated in animal models and humans (4Davis M. The role of the amygdala in conditioned and unconditioned fear and anxiety.in: Aggleton J.P. The Amygdala. Oxford University Press, Oxford, UK2000: 213-288Google Scholar, 5LeDoux J.E. The amygdala.Curr Biol. 2007; 17: 868-874Abstract Full Text Full Text PDF PubMed Scopus (1031) Google Scholar, 8Delgado M.R. Olsson A. Phelps E.A. Extending animal models of fear conditioning to humans.Biol Psychology. 2006; 73: 39-48Crossref PubMed Scopus (287) Google Scholar) and highlights the central role of the amygdala in fear acquisition, storage, and expression. The amygdala is thought to be the site of association and memory storage for simple cued fear conditioning, with projections to the brainstem and hypothalamus mediating autonomic fear expression and projections to the ventral striatum mediating the use of actions to cope with fear (5LeDoux J.E. The amygdala.Curr Biol. 2007; 17: 868-874Abstract Full Text Full Text PDF PubMed Scopus (1031) Google Scholar). In addition, the hippocampus plays an important role in contextual modulation of fear, supporting the acquisition of fears to contexts and guiding the contextually dependent expression of fear (9Fanselow M.S. Contextual fear, gestalt memories, and the hippocampus.Behav Brain Res. 2000; 110: 73-81Crossref PubMed Scopus (604) Google Scholar). Although their specific roles are less clearly defined, the insula and dorsal anterior cingulate cortex are also proposed to modulate fear acquisition (10Shi C. Davis M. Pain pathways involved in fear conditioning measured with fear-potentiated startle: Lesion studies.J Neurosci. 1999; 19: 420-430PubMed Google Scholar, 11Milad M.R. Quirk G.J. Pitman R.K. Orr S.P. Fischl B. Rauch S.L. A role for the human dorsal anterior cingulate cortex in fear expression.Biol Psychiatry. 2007; 62: 1191-1194Abstract Full Text Full Text PDF PubMed Scopus (378) Google Scholar). The inhibition or control of conditioned fear requires the ventromedial prefrontal cortex (vmPFC), which is necessary for the storage of extinction memory. During extinction retrieval, projections from the vmPFC to inhibitory interneurons within the amygdala diminish fear expression. After extinction, contextual information modulates the competition between the original fear memory and the new extinction memory (7Bouton M.E. Context and behavioral processes in extinction.Learn Mem. 2004; 11: 485-494Crossref PubMed Scopus (1394) Google Scholar). Projections from the hippocampus to the vmPFC and the amygdala appear to mediate this context-dependent expression of extinction (9Fanselow M.S. Contextual fear, gestalt memories, and the hippocampus.Behav Brain Res. 2000; 110: 73-81Crossref PubMed Scopus (604) Google Scholar, 12Ji J. Maren S. Electrolytic lesions of the dorsal hippocampus disrupt renewal of conditional fear after extinction.Learn Mem. 2005; 12: 270-276Crossref PubMed Scopus (136) Google Scholar). During the intentional, cognitive regulation of fear (and negative affect more generally) amygdala activation typically decreases, driven by increased activation of the dorsolateral prefrontal cortex (dlPFC) that, in turn, recruits the vmPFC–amygdala inhibitory pathway that mediates extinction retrieval (6Hartley C.A. Phelps E.A. Changing fear: The neurocircuitry of emotion regulation.Neuropsychopharmacology. 2010; 35: 136-146Crossref PubMed Scopus (369) Google Scholar, 13Ochsner K.N. Gross J.J. The cognitive control of emotion.Trends Cogn Sci. 2005; 9: 242-249Abstract Full Text Full Text PDF PubMed Scopus (3172) Google Scholar, 14Schiller D. Delgado M.R. Overlapping neural systems mediating extinction reversal and regulation of fear.Trends Cogn Sci. 2010; 14: 268-276Abstract Full Text Full Text PDF PubMed Scopus (225) Google Scholar). In short, the amygdala, the vmPFC, and the hippocampus collectively support the acquisition, storage, retrieval, and contextual modulation of fear acquisition and extinction (5LeDoux J.E. The amygdala.Curr Biol. 2007; 17: 868-874Abstract Full Text Full Text PDF PubMed Scopus (1031) Google Scholar, 15Quirk G.J. Mueller D. Neural mechanisms of extinction learning and retrieval.Neuropsychopharmacology. 2008; 33: 56-72Crossref PubMed Scopus (1284) Google Scholar). Although anxiety can be distinguished from fear in a number of important respects (2Sylvers P. Laprarie J. Lilienfeld S. Differences between trait fear and trait anxiety: Implications for psychopathology.Clin Psychol Rev. 2011; 31: 122-137Crossref PubMed Scopus (242) Google Scholar), several prominent theories propose that dysregulation of the neurocircuitry implicated in the acquisition and modulation of conditioned fear may be critically involved in the etiology and maintenance of anxiety (16Rachman S. Neo-conditioning and the classical theory of fear acquisition.Clin Psychol Rev. 1991; 11: 155-173Crossref Scopus (171) Google Scholar, 17Bouton M.E. Mineka S. Barlow D.H. A modern learning theory perspective on the etiology of panic disorder.Psychol Rev. 2001; 108: 4-32Crossref PubMed Scopus (820) Google Scholar, 18Mineka S. Zinbarg R. A contemporary learning theory perspective on the etiology of anxiety disorders—It's not what you thought it was.Am Psychol. 2006; 61: 10-26Crossref PubMed Scopus (687) Google Scholar). 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Overgeneralization of conditioned fear as a pathogenic marker of panic disorder.Am J Psychiatry. 2010; 167: 47-55Crossref PubMed Scopus (420) Google Scholar). Additional brain regions may contribute to differences in emotional expression and awareness associated with anxiety. Although the amygdala clearly mediates cue-evoked phasic fear responses to threat-related stimuli, the bed nucleus of the stria terminalis, a region in the ventral basal forebrain referred to as part of the “extended amygdala,” appears to support a more sustained state of arousal and vigilance characteristic of anxious individuals (3Davis M. Walker D.L. Miles L. Grillon C. Phasic vs sustained fear in rats and humans: role of the extended amygdala in fear vs anxiety.Neuropsychopharmacology. 2010; 35: 105-135Crossref PubMed Scopus (1069) Google Scholar, 30Somerville L.H. Whalen P.J. Kelley W.M. 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Oxford University Press, Oxford, UK2000: 213-288Google Scholar, 5LeDoux J.E. The amygdala.Curr Biol. 2007; 17: 868-874Abstract Full Text Full Text PDF PubMed Scopus (1031) Google Scholar, 6Hartley C.A. Phelps E.A. Changing fear: The neurocircuitry of emotion regulation.Neuropsychopharmacology. 2010; 35: 136-146Crossref PubMed Scopus (369) Google Scholar, 7Bouton M.E. Context and behavioral processes in extinction.Learn Mem. 2004; 11: 485-494Crossref PubMed Scopus (1394) Google Scholar). Although precisely how these shared networks jointly contribute to anxiety and decision-making is unclear, this overlap suggests the brain systems mediating fear and anxiety are intertwined with those underlying the computation of value and choice. 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